Volume 31(3): 1–380

METAMORPHOSIS ISSN 1018–6490 (PRINT) ISSN 2307–5031 (ONLINE) LEPIDOPTERISTS’ SOCIETY OF

An overview of -host-parasitoid associations for southern Africa, including an illustrated report on 2 370 African Lepidoptera-host and 119 parasitoid-Lepidoptera associations Published online: 3 November 2020

Hermann S. Staude1*, Marion Maclean1, Silvia Mecenero1,2, Rudolph J. Pretorius3, Rolf G. Oberprieler4, Simon van Noort5, Allison Sharp1, Ian Sharp1, Julio Balona1, Suncana Bradley1, Magriet Brink1, Andrew S. Morton1, Magda J. Botha1, Steve C. Collins1,6, Quartus Grobler1, David A. Edge1, Mark C. Williams1 and Pasi Sihvonen7 1Caterpillar Rearing Group (CRG), LepSoc Africa. [email protected], [email protected], [email protected], [email protected], [email protected], [email protected], [email protected], [email protected], [email protected], [email protected], [email protected], [email protected] 2Centre for Statistics in Ecology, Environment and Conservation, Department of Statistical Sciences, University of Cape Town, South Africa. [email protected] 3Department of Agriculture, Faculty of Health and Environmental Science. Central University of Technology, Free State, Bloemfontein, South Africa. [email protected] 4CSIRO National Collection, G. P. O. Box 1700, Canberra, ACT 2701, Australia. [email protected] 5Research & Exhibitions Department, South African Museum, Iziko Museums of South Africa, Cape Town, South Africa and Department of Biological Sciences, University of Cape Town, Rondebosch, South Africa. [email protected] 6African Research Institute (ABRI), Nairobi, Kenya. [email protected] 7Finnish Museum of Natural History, Helsinki, Finland. [email protected] * Corresponding author

Copyright © Lepidopterists’ Society of Africa

Abstract: We present an overview of the known host associations of larval Lepidoptera for southern Africa, based on a database of 11 628 rearings, including all Caterpillar Rearing Group (CRG) records and other published records. Rearings per Lepidoptera family show some bias in the rearing effort towards the more conspicuous families, ectophagous groups and non-detritus-feeders but in general follow diversity. Recorded Lepidoptera host associations per host family for southern Africa are shown. Data analyses revealed the following general trends: of the 20 most reared species 13 are polyphagous; are the most utilised family with 2 122 associations, followed by Asteraceae (600), Malvaceae (564) and Anacardiaceae (476); 98.8 % of hosts are vascular ; and of the 19 most utilised host species 18 are common trees or shrubs. We discuss possible reasons behind these trends, particularly the high utilisation of Fabaceae and the widespread use of trees and shrubs as hosts. We compare recorded host species numbers with species diversity for the 19 most recorded host families and discuss possible reasons for the low utilisation of four plant families with an exceptionally low percentage of Lepidoptera host species / plant host species diversity. All Lepidoptera families for which more than 100 rearings have been recorded (21 families) utilise one (or two in the case of Pyralidae, Nolidae and Hesperiidae) plant family exponentially more than any of the other families, with resulting histograms forming hyperbolic curves, as are typical of distributions of taxonomic assemblages in nature. We calculate an exponential factor to quantify this phenomenon and show that for all 21 Lepidoptera families one host family is utilised 6–33 times more than the average use of other host families. In this paper, the larvae and adults of 953 African, mostly South African, Lepidoptera species reared by the CRG between January 2016 and June 2019 are illustrated together with pertinent host information. 119 Lepidoptera-parasitoid associations are reported, comprising seven hymenopteran families and one dipteran family. With the current data release, larval host association records are now available for 2 826 Lepidoptera species in the southern African subregion, covering about 25 % of the described fauna.

Key words: Lepidoptera, , Diptera, caterpillar, larva, , butterfly, South Africa, Africa, Afrotropical region, pupa, host, life-history, host-use, host-specialisation, parasitoid.

Citation: Staude, H.S., Maclean, M., Mecenero, S., Pretorius, R.J., Oberprieler, R.G., Van Noort, S., Sharp, A., Sharp, I., Balona, J., Bradley, S., Brink, M., Morton, A.S., Botha, M.J., Collins, S.C., Grobler, Q., Edge, D.A., Williams, M.C. & Sihvonen, P. (2020). An overview of Lepidoptera-host-parasitoid associations for southern Africa, including an illustrated report on 2 370 African Lepidoptera-host and 119 parasitoid-Lepidoptera associations. Metamorphosis 31(3): 1–380.

INTRODUCTION Plants, insect herbivores and their parasitoids and predators constitute over 75 % of all multicellular diversity of terrestrial life on earth (Price, 2002). All three Received: 15 August 2020 are crucial for ecosystem functioning. The effect of Published: 3 November 2020 disruption of the ecological balance at these trophic levels Copyright: This work is licensed under the Creative Commons can be clearly seen in the devastating impact of alien Attribution-NonCommercial-NoDerivs 3.0 Unported License. To invasive species across the globe (Vilá et al., 2011). Yet view a copy of this license, send a letter to Creative Commons, Second Street, Suite 300, San Francisco, California, 94105, USA, even fundamental knowledge on ecological interactions at or visit: http://creative commons.org/licenses/by-nc-nd/3.0/ these three trophic levels for the Afrotropical region is largely lacking.

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A compilation of Lepidoptera-host associations recorded not included in the previous papers for various reasons for southern Africa (including all published records to (e.g. late submission to the project). date) conducted in 2012 registered such associations for less than 7 % of the Lepidoptera species in southern Comprehensive details of the methods employed by the Africa and for only a handful of Lepidoptera-parasitoid CRG were published in Staude et al. (2016a), which associations (Staude & Kroon, 2012). To address this remain the same for this publication (unless otherwise problem, at least for Lepidoptera-host-parasitoid stated below), and only a shortened version is presented interactions, the Lepidopterists’ Society of Africa here. (LepSoc Africa) launched a successful citizen science project called the Caterpillar Rearing Group (CRG), Terminology details of which can be found in Staude et al. (2016a). The The terms ‘caterpillar’ and ‘larva’ are used CRG compiles selected rearing data into master lists, each interchangeably in this article, although they are not fully representing a major Lepidoptera taxonomic group. The synonymous. ‘Larva’ (plural: larvae) is the technical term main purpose of the master lists is to create a vehicle by for the life stage of any holometabolous insect between which this large and growing set of life-history data can egg and pupa, whereas ‘caterpillar’ is a vernacular term become accessible to other users, from scientists to for the (generally free-living) larvae of and conservationists, naturalists and other parties. (Lepidoptera) as well as those of sawflies

Parasitoids are defined by their behaviour of ultimately (Hymenoptera). The rearing efforts of the CRG do not killing their hosts (Price, 1975; Godfray & Godfray, 1994; cover the caterpillars of sawflies, however.

Mills, 2009; Quicke, 2015). It is usually the larval stage of The term ‘host’ is relative and depends on the context in the parasitoid that attacks the host invertebrate (usually which it is being used in the text. A Lepidoptera ‘host’ the immature stages), whereas the adults are free-living refers to the source organism of the food used by the (Vinson, 1976; Godfray & Godfray, 1994). In contrast to Lepidoptera larva. This is usually a living plant, but it may true parasites, which do not kill the host, parasitoids only also be other living organisms or their detritus. A require a single host to develop into adults (Price, 1975; parasitoid ‘host’ is the lepidopteran taxon from which the Godfray & Godfray, 1994). In the class Insecta, several parasitoid was reared, generally its egg, larval or pupal orders are known to include parasitoid species stage. Parasitoids also use many other organisms as hosts (Coleoptera, Diptera, Hymenoptera, Lepidoptera, but in the context of this paper it refers to Lepidoptera. Neuroptera, Strepsiptera and Trichoptera), with ca. 10 % of described insect species being parasitoids (Price, 1975; The term ‘rearing’ refers to a rearing event in which a Mills, 2009). This is the first time that we report on larval host association was established by observation that parasitoid-Lepidoptera associations. Although many more the larva was feeding on the host. The rearing of multiple parasitoids have been reared by CRG members, only 119 conspecific caterpillars from the same host and same parasitoid-Lepidoptera associations have been processed locality at the same time constitutes a single rearing. and are dealt with in a preliminary manner here at family When a single lava was reared on multiple host species, level, prior to undergoing further identification of the these were treated as separate rearings, with the host on parasitoid species. which the larva was found in the field regarded to represent a natural host association and the others as The first CRG publication reported on 1 778 rearings established only in captivity. comprising 962 species of Afrotropical Lepidoptera (Staude et al., 2016a, 2016b). The second CRG The term ‘master lists’ refers to lists compiled by the CRG publication reported on another 458 rearings comprising containing pertinent data for each rearing received from 424 taxa of Afrotropical Papilionoidea (Congdon et al., members, including: unique rearing number; Lepidoptera 2017). We present master lists on a further 2 370 rearings, species, subfamily and family; host species and family; comprising 953 species of Afrotropical Lepidoptera and locality; collecting date, pupation date, eclosion date; adding an additional 641 species to the total number rearer name; photograph of final-instar larva; photograph reared, thus bringing the total of Lepidoptera represented of adult. in the CRG master lists to 2 027 species. The term ‘main database’ refers to a database of Kroon (1999) published lists for Lepidoptera-host Afrotropical Lepidoptera-host associations kept by the associations known at the time, but no overview of the first author, which includes all CRG records, all records state of knowledge of Lepidoptera-host associations for published by Kroon (1999), records extracted from southern Africa had been published before. We here specimen label data and other published records (Fig. 1). present such an overview of our knowledge of There is a wealth of additional published rearing Lepidoptera-host associations for southern Africa based information available for the Afrotropical Region outside on the rearing efforts of CRG members as well as other of the southern Africa subregion, but most of this has not published Lepidoptera-host associations included in our yet been included in the database. The current database, main database of such associations. We briefly discuss however, is deemed to be a comprehensive list of the new insights emerging from this dataset. known Lepidoptera larval host associations for the southern Africa subregion. The term ‘southern Africa’ MATERIAL AND METHODS comprises South Africa, Lesotho, eSwatini, Botswana, Namibia, Zimbabwe and Mozambique south of the This paper reports on rearings made by CRG members Zambezi river. during the period of January 2016 – June 2019, as well as on rearings conducted at an earlier date but which were

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6000 These were checked as far as possible by M. Maclean. 5500 5491 Many vascular plants that were difficult to identify were 5000 posted on iNaturalist (www.inaturalist.org), where they were identified or their identification confirmed by 4000 experts. Dried herbarium specimens were kept in many cases where plants could not be identified. 3000 For parasitoids, the CRG management team is currently in 2000 the process of preparing voucher specimens for submission to taxon specialists for identification. The No. of rearing records rearing of No. 1000 619 18 Hymenoptera will be photographed at high resolution at 0 the Iziko South African Museum and their data added to CRG Kroon Other Specimen WaspWeb (www.waspweb.org), the online resource for (1999) published label data information on , and ants of the Afrotropical literature biogeographical region. So far we have not found a taxon Source specialist for African dipteran parasitoids.

Figure 1 ‒ Source of rearing records in the main database of The classification of the Lepidoptera largely follows African Lepidoptera-host associations (n = 11 628). Nieukerken et al. (2011) and Krüger (2020). Data in the tables regarding Lepidoptera species diversity for Approach and protocol southern Africa were similarly extracted from Krüger (2020). Members of the public were and still are invited to rear the larvae of African Lepidoptera and keep records of the Plant names and classification follow the portal Plants of rearings. They are encouraged and guided to rear eggs and the World Online (www.plantsoftheworldonline.org) for larvae of moths or butterflies they come across to the vascular plants (Spermatophyta), in collaboration with adulthood, to photograph all life stages and hosts as well the South African National Biodiversity Institute as parasitoids, to record all observations, to preserve (SANBI). For the non-vascular plants and fungi, we do not voucher specimens of the reared moth or butterfly as well have specific authorities, taxon specialists or voucher as of the host and any parasitoids, and to submit the specimens, and most of these remain identified only at records and vouchers to the CRG management team. The higher ranks. following minimum information is required with each submission, generally by e-mail to the CRG management: The localities recorded in the master lists refer to the place locality description including longitude and latitude, name of collection of the specimen, from which the rearing of host if available, date of collection of life stage, date of commenced, be it an early stage (usually egg or larva) or pupation, date of emergence of adult, photographs of an adult female that laid eggs. caterpillar, pupa, adult, host and parasitoid. Notes on interesting observations may also be submitted. All Blank cells in the master lists mean that we have been photographs and data submitted are electronically stored unable to identify the specimen to the denoted taxon level. by CRG management under the names of the individual The tables and information presented in the overview of contributors. At the time of writing, more than 66 500 Lepidoptera-host associations for southern Africa are photographs and documents have been received for based on the main database. safekeeping.

Storage of submitted material Collating the rearing records into master lists CRG members are encouraged to preserve and submit The main current output of the CRG is the master lists, voucher specimens of reared pupae, adults, hosts and comprising rearing records of Afrotropical Lepidoptera parasitoids to the project team. Adult Lepidoptera submitted by CRG members. specimens, when received, are mounted and labelled in

Identifications of Lepidoptera and parasitoid taxa are the standard way for Lepidoptera (Richardson, 2015) and based as far as possible on available voucher specimens. initially stored in the Staude Collection (private collection Bombycoidea are identified or confirmed by R.G. of H.S. Staude housed in Magaliesburg, South Africa). Oberprieler and all other Lepidoptera are identified, or Subsequently, the specimens are, when possible, donated rearer identifications confirmed, by H. S. Staude, unless to taxon specialists for confirmation of species otherwise stated in the master lists where determinations identifications in the master lists and for further were made by taxon specialists. Hymenoptera are taxonomic work. identified by Simon van Noort unless otherwise stated. Parasitoids are initially stored in ethanol vials in the The parasitoid wasps covered in this article were Staude Collection. The Hymenoptera are then transferred identified based on photographs submitted by the rearers to the Iziko South African Museum for identification and but will be verified once the voucher specimens have been permanent storage. The Diptera are transferred to Johan examined. Pretorius who will distribute them to taxon specialists for

Identification of host species are mostly provided by the identification when such have been found. individual rearers using the many excellent illustrated field guides that are available for South African plants.

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Pressed plant specimens received are similarly initially parasitoids (Tables 2 and 3). kept in the Staude collection until a more suitable herbarium depository for them can be found. It is expected that parasitoids could attack all lepidopteran species, but a sustained rearing effort would be required RESULTS & DISCUSSION to eventually obtain a more comprehensive record of parasitoidism. Hundreds of parasitoids have already been During the period of this report (January 2016 – June reared by CRG members, but these still need to be 2019), 2 370 submissions comprising 953 Lepidoptera identified and processed. species were made by 59 CRG members (individuals or teams) and processed by the CRG management. These are A brief overview of the state of knowledge on presented here as 28 CRG master lists (pages 21‒380) and Lepidoptera-host associations for southern Africa should be regarded as an addition to the CRG master lists based on the main database published by Staude et al. (2016a,b) and Congdon et al. (2017). The contributions made by each member or team Establishing host associations for Afrotropical for this latest batch are shown in Table 1 (page 11). The Lepidoptera is a huge task considering the size of the region and the incredible diversity of Lepidoptera and explanation to the master lists is given on page 20. vascular plants that the region contains. The database is The total CRG master lists currently contain 4 524 valid overwhelmingly biased towards the southern Africa rearings comprising 3 275 different host-associations for subregion (Table 4: page 13). Figure 2 shows a map of 2 027 putative species of Afrotropical Lepidoptera. Host rearing localities (where location data are available). This associations have been recorded for the first time for at bias is due to the fact that most CRG members reside in least 652 species, not including the many specimens that southern Africa, but also because most of the host could not be identified to species level, often due to a lack associations available in the published literature for the of expertise but mostly due to an unresolved . rest of the Afrotropical region have not yet been captured in our database. The following results, therefore, are an The main database contains records of 11 628 rearings of attempt at providing a brief overview of our current Lepidoptera-host associations. For southern Africa, knowledge for southern Africa only, based on the main Lepidoptera-host associations are now available for some database. 2 815 putative species, comprising ~25 % of the described Lepidoptera fauna (10 839 species), effectively more than doubling the proportion of about 7 % estimated in 2012 (Staude & Kroon, 2012).

CRG parasitoid rearing submissions

To date, we have processed 119 parasitoid rearing submissions. Of these, 27 % (n = 32) are records of true flies (Diptera) and 73 % (n = 87) are records of wasps (Hymenoptera). The high number of Hymenoptera submissions comes as no surprise, as this order contains ca. 77 % of all known insect parasitoids (Mills, 2009).

The most important families of hymenopteran parasitoids are Braconidae, Chalcididae and (Quicke, 2015). From the records reviewed thus far, seven hymenopteran families were found to parasitise 14 lepidopteran families (Table 2: page 12). Of these, about a)

23 lepidopteran species and three hymenopteran species have been identified so far.

For the dipteran parasitoids, the family Tachinidae appears to represent all submissions received, pending further confirmation by taxon specialists. The family constitutes one of the largest families of Diptera, and all known species are endoparasites of (and sometimes of centipedes) (Barraclough & Londt, 2008). From records received to date, eight lepidopteran families were parasitised by flies (Table 3: page 13). Of these, 14 lepidopteran species have been identified.

Six lepidopteran families (, Geometridae, b) Lasiocampidae, Noctuidae, Notodontidae and Pieridae) acted as hosts for both flies and wasps, and seven subfamilies (Ennominae, Erebinae, Heliothinae, Figure 2 – Maps of a) the Afrotropical region and b) Lasiocampinae, , Pierinae and Plusiinae) southern Africa, depicting locations (where known) of were common hosts of both dipteran and hymenopteran recorded Lepidoptera-host associations (n = 4 478 rearings).

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Host associations for southern Africa were recorded for identified to at least the genus level. Only 128 (1.2 %) of 66 Lepidoptera families (Table 5: page 14). For 24 the 10 981 host associations recorded were for non-plant rearings the Lepidoptera family could not be identified species, confirming that the vast majority of Lepidoptera and for 565 rearings specimens could not be identified to are indeed phytophagous. Fabaceae is by far the family of species level. The main database now has records for vascular plants most utilised by southern African 2 826 identified Lepidoptera species, of which 1 439 are Lepidoptera, with 2 122 associations recorded. The next from the CRG. most-utilised families are Asteraceae (600), Malvaceae (564) and Anacardiaceae (476). Table 6 compares the number of recorded host associations to species diversity for the most represented The 19 plant species most commonly utilised by 19 Lepidoptera families for southern Africa. As can be Lepidoptera are shown in Table 9 (page 17). The three expected, the well-known showy families attracted a high most frequently used host species are: karroo percentage of rearing effort compared to species diversity (Fabaceae) with 356 associations of 158 Lepidoptera (Papilionidae 623 % and 642 %), whereas the species; Gymnosporia buxifolia (Celastraceae) with 144 less showy families showed a much lower percentage associations of 67 species; and Ziziphus mucronata (Gelechiidae 27 % and Pyralidae 34 %). The relatively (Rhamnaceae) with 115 associations of 48 species. All high percentage for Tortricidae (79 %) is probably due to three of these species are common trees that are the poorly worked taxonomy for the family in southern widespread across southern Africa. The high utilisation of Africa, with many known undescribed species. the Australian wattle mearnsii is probably an indication that the secondary defensive metabolites Table 6 ‒ The number of Lepidoptera-host associations produced by this species are not very different from those (rearings) per Lepidoptera family for southern Africa compared produced by the closely related local Acacieae, making to the number of species comprising each Lepidopteran family the host shift to this exotic species rather easy. This exotic (species diversity) for the 19 Lepidoptera families with the most species is also now common and widely distributed number of rearings (n = 10 106). throughout southern Africa.

No. of No. of % rearings/ Family rearings species species With the exception of Ehrharta erecta (Poaceae), all the vascular plants on this list are widespread common shrubs Erebidae 1 769 1 565 113 or trees. All the records for E. erecta are for Hesperiidae Geometridae 1 621 1 531 106 and . It is probable that this plant is on the Lycaenidae 1 063 509 208 list of the 20 most utilised vascular plants because of Noctuidae 972 967 101 rearings from eggs under artificial conditions by Lepidopterist specialists, rather than representing natural Nymphalidae 852 324 263 host associations, or it may be due to erroneous reporting Saturniidae 520 81 642 in the literature, where the text “Ehrharta erecta and other 511 111 460 grasses” is repeatedly used for most grass-feeding species.

Lasiocampidae 486 227 214 The data suggest that widespread trees and shrubs with a Hesperiidae 339 155 219 large individual biomass are particularly prone to being Crambidae 296 526 56 used by Lepidoptera as hosts, but this could also be a case of the under-sampling by caterpillar rearers on herbaceous Pieridae 286 64 447 vascular plants, which are often more difficult to sample. Notodontidae 246 225 110 Table 10 (page 17) compares recorded host associations Tortricidae 236 298 79 and ‘recorded host species’ to ‘host species diversity’ for Pyralidae 215 630 34 the 19 most utilised host families for southern Africa. In Gelechiidae 160 597 27 this comparison the percentage of associations per species Gracillariidae 159 175 91 diversity for Fabaceae was rather low (34 %, comprising only 5.6 % of the flora), indicating that many Fabaceae Nolidae 149 159 94 species have probably not yet been sampled for Papilionidae 137 22 623 Lepidoptera herbivory, possibly because of the many Nepticulidae 89 125 71 localised herbaceous species in this family. This is the case as well for the other plant families on the list, The 20 most commonly reared Lepidoptera species are providing an indication of how much there is to do and shown in Table 7 (page 15). The African Bollworm, that under-sampling is still a major problem. Helicoverpa armigera, recorded from 136 host-plant species belonging to 39 plant families, is by far the most Plant families with an exceptionally low percentage are encountered species. Eleven of the 20 species on this list highlighted with an asterisk (Table 10: page 17). For utilise 10 or more host families. By contrast, Chiasmia Apocynaceae (8 %) and Euphorbiaceae (15 %) the reason brongusaria and Cabera strigata utilise only one host for the low utilisation by Lepidoptera can probably be ascribed to their known numerous well developed family. defensive secondary metabolites (see Livishultz et al. Recorded Lepidoptera-host associations per host taxon (2018) for Apocynaceae; and Mwine & Van Damme (mostly family) for the southern Africa are shown in Table (2011) for Euphorbiaceae). 8 (page 15). Host associations were recorded for 174 families. Over 2 600 host taxa could be Poaceae (13 %) are well known for their structural

Staude et al. / Metamorphosis 31(3): 1–380 6 defensive mechanisms, incorporating abrasive silicates Table 11 ‒ Recorded Geometridae-host associations per host into their leaf cells (Kaufman et al., 1985), which seems plant family, for the 27 most used families (n = 1 380). to prevent their utilisation as food by major Lepidoptera No. of No. of groups and probably contributes to the low percentage Plant family Plant family found here. For instance, there are no records of any rearings rearings Geometridae (the second-most diverse Lepidoptera Fabaceae 438 Santalaceae 17 family) utilising Poaceae as hosts in our database (Staude Celastraceae 184 Icacinaceae 14 et al., 2019). Anacardiaceae 100 Primulaceae 14 A contributing factor in Africa could be the grazing Rhamnaceae 97 Sapindaceae 14 pressure on Poaceae from large mammals. Caterpillars run Asteraceae 95 Stilbaceae 13 a high risk of being eaten by these grazers and appear to need particular defensive strategies to be able to feed on Rubiaceae 76 Ochnaceae 12 grasses, such as having urticating spines (as in the Proteaceae 46 Phyllanthaceae 12 saturniid Decachorda, the lymantriine Psalis africana and Combretaceae 36 Rosaceae 12 some Eupterotidae) or being large and spiny and feeding only on large grasses (the saturniid Bunaeopsis) or feeding Ericaceae 35 Asparagaceae 11 only at night (many Eupterotidae and Noctuidae) or Zamiaceae 28 Ebenaceae 11 feeding low down near the root base where grazers do not Ranunculaceae 26 Myrtaceae 11 reach (Crambinae). Oleaceae 20 Plumbaginaceae 11 Some collecting bias should also not be disregarded, as no Loranthaceae 19 Stangeriaceae 11 rearings are recorded for Crambinae in the data, which are well known to feed on grasses in other regions. Euphorbiaceae 17

The low percentage for Asteraceae (16 %) is more Table 12 ‒ Recorded Pieridae-host associations per host plant difficult to explain and may be an artefact of under- family (n = 285). sampling as most vascular plants in this family are low- No. of No. of growing herbs that make it difficult to find caterpillars. It Plant family Plant family is expected that a concerted effort by CRG members to rearings rearings look for caterpillars on Asteraceae will reveal many Capparaceae 159 Lauraceae 1 interesting new associations. Fabaceae 37 Moraceae 1

The highest percentage (115 %) is for Loranthaceae and Brassicaceae 30 Olacaceae 1 reflects the extensive work done in the past on the life Loranthaceae 13 Oleaceae 1 histories of the Theclinae (Lycaenidae) (Congdon et al., 2017), which represents a single radiation, and this work Salvadoraceae 11 Phyllanthaceae 1 has evidently inflated the data abnormally. Santalaceae 6 Polygonaceae 1

A further interesting result is that most Lepidoptera Celastraceae 5 Resedaceae 1 families seem to utilise one (or two in the case of Clusiaceae 4 Rhizophoraceae 1 Pyralidae, Nolidae and Hesperiidae) plant family exponentially more than any of the other families. For Poaceae 4 Rosaceae 1 example, Geometridae use 91 plant families as hosts but Apocynaceae 2 Sapindaceae 1 use Fabaceae exponentially more than any other family Asteraceae 2 Tropaeolaceae 1 (Table 11), and Pieridae utilise 23 plant families but use Capparaceae exponentially more than the rest (Table 12). Euphorbiaceae 1

In order to quantify this phenomenon, of a host family The fact that this pattern applies to all 21 Lepidoptera being utilised exponentially more than others, we families and is derived from a dataset of 11 055 records calculated an exponential factor (A) using the formula A accumulated across the subcontinent over many years by = D / (B - D) x E, where B = records for Lepidoptera more than 150 rearers, who were not subjected to any family, D = records for host family, E = host families standardised sampling method, suggests that this recorded for Lepidoptera family. Table 13 (page 18) phenomenon is not an artefact of sampling bias but real. shows this exponential factor for all the Lepidoptera When these host usages are plotted in numerical families in southern Africa for which our dataset contains sequences (Figs 3ab, 4), the resulting histograms form more than 100 rearing records (L100). This exponential hyperbolic or ‘hollow’ curves, as are typical of factor quantifies the exponential nature of host family distributions of taxonomic assemblages in nature (e.g. usage for a given Lepidoptera family. For instance, in Willis & Udny Yule, 1922; Dial & Marzluff, 1989; Erebidae, Fabaceae are used 33 times more than the Scotland & Saunderson, 2004; Richardson & Oberprieler, average use of the their other host families. 2007). This common, non-random distribution is evidently a true and universal evolutionary pattern that appears to reflect the fractal geometry of taxic diversity

(Burlando, 1990) and seems largely related to the division

Staude et al. / Metamorphosis 31(3): 1–380 7

500 a) 450 400 350 300 250 200

No. of rearings 150 100 50 0

Plant host family

160 b) 140 120 100 80 60 No. of rearings 40 20 0

Plant host family

Figure 3 ‒ Graph of a) Geometridae-host associations (rearings) for all 91 host families (n = 1 597 rearings) and b) Pieridae-host associations per plant family (using data in Table 12; n = 285 rearings), showing the hollow curve phenomenon for a family that utilises one host family exponentially more than others.

120

100

80

60

40 No. of rearings 20

0

Plant host family

Figure 4 ‒ Graph of Hesperiidae-host associations (rearings) per plant family, showing the hollow curve phenomenon for a family that utilises two host families exponentially more than others (n = 336 rearings).

Staude et al. / Metamorphosis 31(3): 1–380 8

of ecological niche space available to diversifying Table 15 ‒ Recorded Lepidoptera-host lineages (Dial & Marzluff, 1989; Richardson & associations per genus in the tribe Oberprieler, 2007). Our data seem to confirm that Acacieae. taxonomic host usage of Lepidoptera follows this same Genus No. of evolutionary pattern. rearings For 12 of the 21 Lepidoptera families (L100), the recorded Vachellia 548 exponentially-used host family is Fabaceae. 747 Senegalia 168 Lepidoptera species have been recorded feeding on 359 Acacia 132 species of Fabaceae in southern Africa (Table 10: page 17). The high species diversity of Fabaceae in southern Undetermined to genus 81 Africa (6 374) may possibly explain the high utilisation of Total 929 this family by Lepidoptera for the region, but this is unlikely because the recorded hosts comprise just 5.6 % It would be interesting to compare host use of of the diversity of the family in the region (Table 10: page exponentially used host families in terms of biomass 17). rather than species diversity, if such figures were available. Equally useful would be a comparison between A more likely factor for this pattern is the prominence of host use and the secondary defensive metabolites these the family in terms of its common and widespread vascular plants produce. presence and large biomass across a large part of the southern African landscape (e.g. tribe Acacieae), coupled CONCLUSION with the fact that Fabaceae may be more nutritious due to their well-known symbiosis with nitrogen-fixing , This brief overview of Lepidoptera-host associations has an important property in a generally nutrient-poor region. produced more questions than answers, but it is hoped that Edge & van Hamburg (2009) reviewed the frequent use of this will stimulate further in-depth investigations into the Fabaceae in myrmecophilous lycaenid larvae and found intricate relationships between Lepidoptera and their evidence of high content of essential amino acids. Many hosts. Poaceae also have associations with nitrogen-fixing bacteria, but the abrasive silicates they incorporate into It is clear from the summaries of our data presented here their leaf structure and other ecological factors probably that the distribution of host plant taxa among Lepidoptera prevent many Lepidoptera from utilising them (as needs to be investigated further, and further analyses are discussed above). required to corroborate the patterns found here. This should include the addition of host records from other The usage of hosts within the Fabaceae by Lepidoptera is sources from other regions in Africa (e.g. the literature, unevenly spread across the different lineages (Table 14: notwithstanding its unreliability in many cases) as well as page 18). Of the 2 187 rearing records for the family, 929 the more robust delimitation of lepidopteran taxa by (43 %) refer to the tribe Acacieae alone, which is means of phylogenetic methods (many are not significant seeing that these records span only 5.6 % of the demonstrably monophyletic, especially at the tribal level Fabaceae species in the region (Table 10: page 17). Even and below). Phylogenetic studies should also be able to though much more extensive sampling of Lepidoptera- elucidate the evolutionary history of host usage and host Fabaceae associations is required before definitive shifts (from ancestral to secondary hosts). It is our conclusions can be reached about this pattern, it appears intention to incorporate the wealth of new information that that the widespread utilisation of Acacieae is the main modern molecular phylogenomics can bring in this regard factor in the predominance of Fabaceae as larval hosts into our database so as to enable further analysis of host among the Lepidoptera fauna of southern Africa. This is patterns. This should be especially revealing once the unlikely to change radically when more data become phylogenetic positions of both the Lepidoptera and their available. Host usage is also uneven among the genera in hosts become better resolved between the family and the tribe Acacieae (Table 15), in that 59 % of the recorded genus levels. host associations apply to the genus Vachellia, and this ratio increases to 69 % when the unnatural associations The CRG has now entered its eighth year and has achieved with the exotic genus Acacia are excluded. Furthermore, much in the form of new data regarding the life-histories host usage is also greatly skewed in Vachellia (Table 16: of southern African Lepidoptera. It has effectively page 19), in that 68 % of the recorded associations are with doubled, in seven years, the amount of Lepidoptera-host one species only, V. karroo, which thus emerges as the associations that had been accumulated in more than a most extensively utilised host by Lepidoptera in southern century before its inception in 2012. For the last few years Africa. The abundance and wide distribution of this now, new host associations are being accumulated at a rate species in the region is probably partly responsible for this of more than two per day by the CRG and it is hoped that pattern, but it may also be more palatable to Lepidoptera this trend will continue in future. larvae in general than other species of the genus, and also than other species of the Acacieae and of Fabaceae The quality of the data is also much improved. CRG overall. Nevertheless, the data show that even at the genus records have many advantages over traditional historical level one species is utilised exponentially more than any host records, which were often derived from ab ovum other and the same ‘hollow” curve is produced (Fig. 5) as rearings in order to obtain good specimens for collections, those at the family level (Figs 3,4).

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400 350 300 250 200 150

No. of rearings 100 50 0

Species

Figure 5 ‒ Graph of Lepidoptera-host associations per species of Vachellia using the data in Table 16, showing the hollow curve phenomenon appearing at genus level. thereby sometimes establishing artificial host- These have greatly enhanced this article. associations. These advantages are: female host choice in the wild is established; parasitoid associations are LITERATURE CITED identified; exact geographical sampling locations are recorded; habitats at locations of sampling are often BARRACLOUGH, D.A. & LONDT, J.G.H. 2008. Order photographed; CRG members are spatially spread across Diptera, pp. 283–321. In: Scholtz, C.H. & Holm, E. nd the region; high-quality photographs of early stages are (Eds) Insects of Southern Africa. 2 ed. Protea, South produced; fresh voucher specimens are frequently Africa. obtained and preserved for further study (including BURLANDO, J. 1990. The fractal dimension of molecular analysis). taxonomic systems. Journal of Theoretical Biology 146(2): 99–114. The resultant improved quality of the data and the rapidly CONGDON, T.C.E., BAMPTON, I. & COLLINS, S.C. accumulating CRG data in the main database will 2017. An illustrated report on the larvae, adults and hopefully continue to enable better analyses and a better host associations of 424 African Lepidoptera taxa understanding of the vital intricate relationships between belonging to the Papilionoidea. A second report of the members of these three fundamental trophic levels upon Caterpillar Rearing Group of LepSoc Africa. which terrestrial life is reliant. Metamorphosis 28: 57–150. DIAL, K.P. & MARZLUFF, J.M. 1989. Nonrandom ACKNOWLEDGEMENTS diversification within taxonomic assemblages. Systematic Zoology 38: 26–37. We thank first and foremost all our fellow caterpillar EDGE, D.A. & VAN HAMBURG, H. 2009. Larval rearers for their contributions, as individually recognised feeding behaviour and myrmecophily of the Brenton in the master lists. Thanks are also due to our taxon Blue, Orachrysops niobe (Trimen) (Lepidoptera: specialists for identifying Lepidoptera in their sphere of Lycaenidae) Journal of Research on the Lepidoptera expertise, who are likewise acknowledged at source. We 42: 21–33. thank the iNaturalist community (www.inaturalist.org) for GODFRAY, H.C.J. & GODFRAY, H.C.J. 1994. supporting the CRG from its inception and for always Parasitoids: behavioral and evolutionary ecology being very helpful when plant identifications are required. (Vol. 67). Princeton University Press. Some of our active participants are from this community. KAUFMAN, P.B., DAYANANDAN, P., FRANKLIN, C. Cape Nature, Ezemvelo KZN Wildlife, Isimangaliso I. & TAKEOKA, Y. 1985. Structure and function of Wetland Park Authority, South African National Parks, silica bodies in the epidermal system of grass shoots. Northern Cape Nature Conservation, Eastern Cape Parks Annals of Botany 55(4): 487–507. and Tourism Agency (ECPTA) and Limpopo Provincial KROON, D.M. 1999. Lepidoptera of Southern Africa, Government are thanked for permission to collect Host-plants & Other Associations – A Catalogue. The caterpillars in areas under their jurisdiction. We thank Lepidopterists’ Society of Africa, 159 pp. Tony Rebelo and Ronel Kloppers for permission to use KRÜGER, M. 2020. Checklist of the Lepidoptera of data contained in their unpublished plant list for southern southern Africa, Metamorphosis 31(2): 1–201. Africa. We are grateful to the two peer reviewers, Konrad LIVSHULTZ, T., KALTENEGGER, E., STRAUB, Fiedler and Axel Hausmann, who have chosen to make S.C.K., WEITEMIER, K., HIRSCH, E., KOVAL, K., their identity known to us, for their very valuable MEMA, L. & LISTON, A. 2018. Evolution of comments and recommendations for future projects. pyrrolizidine alkaloid biosynthesis in Apocynaceae:

Staude et al. / Metamorphosis 31(3): 1–380 10

revisiting the defence de-escalation hypothesis. New mailing_list&mailid=107-moth-challange Phytologist 218: 762–773. doi: 10.1111/nph.15061 STAUDE, H.S., MECENERO, S., OBERPRIELER, R., MILLS, N. 2009. Parasitoids, pp. 748‒750. In: Resh, V.H. SHARP, A., SHARP, I., WILLIAMS, M.C. & & Cardé, R.T. (Eds) Encyclopaedia of Insects. 2nd ed. MACLEAN, M. 2016a. An illustrated report on the Elsevier, U.S.A. larvae and adults of 962 African Lepidoptera species. MWINE, J.T. & VAN DAMME, P. 2011. Why do Results of the Caterpillar Rearing Group: a novel, Euphorbiaceae tick as medicinal plants? A review of collaborative method of rearing and recording Euphorbiaceae family and its medicinal features. lepidopteran life-histories. Metamorphosis 27: 46–59. Journal of Medicinal plants Research 5: 652–662. STAUDE, H.S., MECENERO, S., OBERPRIELER, R., NIEUKERKEN et al. 2011. Order Lepidoptera. In: SHARP, A., SHARP, I., WILLIAMS, M.C. & Zhang, Z.-Q. (Ed.) biodiversity: An outline of MACLEAN, M. 2016b. Supplementary material to: higher-level classification and survey of taxonomic An illustrated report on the larvae and adults of 962 richness. Zootaxa 3148: 212–221. African Lepidoptera species. Results of the Caterpillar PRICE, P.W. 1975. Reproductive strategies of Rearing Group: a novel, collaborative method of parasitoids. In: Evolutionary strategies of parasitic rearing and recording lepidopteran life-histories. insects and mites (pp. 87‒111). Springer, Boston, MA. Metamorphosis 27 Supplement: S1–S330. PRICE, P.W. 2002. Resource-driven terrestrial interaction STAUDE, H.S., SIHVONEN, P. & HAUSMANN, A. webs. Ecological Research 17: 241–247. 2019. A phylogenetic comparison of Afrotropical QUICKE, D.L. 2015. The braconid and ichneumonid Geometridae and their host-plants. In: HAUSMANN parasitoid wasps: biology, systematics, evolution and A. & RAJAEI H. (Eds) Proceedings of the tenth ecology. John Wiley & Sons. Forum Herbulot 2018. Integrative taxonomy, a RICHARDSON, B.J. & OBERPRIELER, R.G. 2007. The multidisciplinary approach to answer complicated diversity of Linnaean communities: a way of detecting taxonomic questions (Stuttgart, Germany, 11‒16 June invertebrate groups at risk of extinction. Journal of 2018). Spixiana 42: 291–320. Insect Conservation 11: 287–297. VILÁ, M. , ESPINAR, J.L., HEJDA, M., HULME, P.E., RICHARDSON, I.D. (Ed.) 2015. Practical Guide to the JAROSˇI´K, V., MARON, J.L., PERGL, J, Study of Lepidoptera in Africa. 2nd edition. SCHAFFNER, U., SUN, Y. & PYSˇEK, P. 2011. Lepidopterists’ Society of Africa. Ecological impacts of invasive alien plants: a meta- http://www.lepsocafrica.org/?p=publications&s=pg analysis of their effects on species, communities and SCOTLAND, R.W. & SANDERSON, M.J. 2004. The ecosystems. Ecology Letters 14: 702–708. significance of few versus many in the tree of life. VINSON, S.B. 1976. Host selection by insect parasitoids. Science 303: 643. Annual review of entomology 21(1): 109‒133. STAUDE, H.S. & KROON, D.M. 2012. The Great WILLIS, J.C. & UDNY YULE, G. 1922. Some statistics Caterpillar Moth Hunt Challenge. Newsletter of the of evolution and geographical distribution in plants Lepidopterists’ Society of Africa. Downloadable from and and their significance. Nature 109: 177– http://www.lepsoc.org.za/oldsite/index.php?option=c 179. om_acymailing&ctrl=archive&task=view&listid=1-

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Table 1 ‒ Valid rearing contributions (n = 2 370 rearings) made by CRG members for the period January 2016 – June 2019.

No. of No. of Rearer/team Rearer/team submissions submissions A. Sharp & I. Sharp 997 A. du Plessis 1 H. S. Staude 573 A. Metcalf 1 J. Balona 126 A. Morton & E. Topp 1 S. Bradley 121 B. Jansen 1 M. Brink 87 C. Barnard 1 M. Maclean 82 C. Huisman 1 A. Morton 70 C. Mateke & M. Imakando 1 M. J. Botha 64 C. Meyer 1 S. Mecenero 62 D. Muller 1 S. C. Collins 27 E. Engelking 1 Q. Grobler 25 F. Visser 1 D. Wood 14 G. Grieve 1 P. Webb 13 H. Heya 1 C. Schuster 10 H. Otto 1 G. Hoile 10 I. Thomas 1 H. Vermaak 7 J. D. Hill 1 P. English 6 J. Grosel 1 P. Vos 6 J. Kemper 1 S. Basel 6 J. Saksida 1 S. du Preez 6 K. W. & G. R. H. Grieve 1 S. E. Woodhall 5 L. Baldwin & A. Morton 1 B. Altenroxel 4 L. Bentley 1 C. Risch 4 L. le Roux 1 J. Groenewald 4 M. Henrion 1 A. Coetzer 2 M. Maclean & M. Brink 1 A. Coetzer & E. Brand 2 M. Olivier 1 A. Konda 2 M. Way 1 C. Reynolds 2 R. Taylor 1 M. Roberts 2 S. Atkins 1 R. Peckover 2

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Table 2 ‒ A list of hymenopteran parasitoids with their lepidopteran hosts, identified to at least family level.

Hymenopteran parasitoid* Lepidopteran host* Family Subfamily Tribe Species Family Subfamily Species Bethylidae ?Tortricidae 1 species NI Braconidae ?Crambidae 1 species NI Braconidae ?Geometridae 1 species NI Braconidae ?Nolidae ?Nolinae 2 species NI Braconidae Erebidae Erebinae 1 species NI Braconidae Geometridae 5 species NI Braconidae Geometridae Ennominae Ascotis reciprocaria Braconidae Geometridae Laurentiinae Haplolabida inaequata Braconidae Geometridae Sterrhinae Palaeaspilates inoffensa Braconidae Noctuidae Noctuinae ?Agrotis segetum Braconidae Noctuidae Noctuinae ?Agrotis sp. Braconidae Noctuidae Noctuinae Agrotis segetum Braconidae Noctuidae Noctuinae Agrotis segetum Braconidae Plutellidae Plutellinae Plutella xylostella Braconidae Tortricidae 1 species NI Braconidae Braconinae Erebidae Boletobiinae Eublemma sp.

Braconidae Braconinae Geometridae Ennominae Isturgia deerraria Braconidae Braconinae ?Archibracon Saturniidae alcinoe servillei Braconidae Euphorinae Erebidae Calpinae Exophyla multistriata Braconidae Microgastrinae ?Crambidae 2 species NI Braconidae Microgastrinae ?Nolidae 1 species NI Braconidae Microgastrinae Erebidae Erebinae robusta Braconidae Microgastrinae Erebidae Lymantriinae 1 species NI Braconidae Microgastrinae Erebidae Lymantriinae Psalis africana Braconidae Microgastrinae Geometridae 3 species NI Braconidae Microgastrinae Geometridae Ennominae Drepanogynis cambogiaria Braconidae Microgastrinae Noctuidae Plusiinae Chrysodeixix chalcites Braconidae Microgastrinae Pieridae Pierinae Pieris brassicae Braconidae Microgastrinae Tortricidae 1 species NI Encyrtidae Noctuidae Plusiinae 1 species NI Eulophidae Entedoninae Noctuidae Plusiinae 1 species NI Eurytomidae Eurytominae Cecidosidae Scyrotis sp. Ichneumonidae ?Nolidae ?Nolinae 1 species NI Ichneumonidae Erebidae Lymantriinae Euproctis bicolor Ichneumonidae Geometridae 4 species NI Ichneumonidae Geometridae Ennominae 1 species NI Ichneumonidae Geometridae Sterrhinae Rhodometra sacraria Ichneumonidae Noctuidae 2 species NI Ichneumonidae Noctuidae Heliothinae Heliothis scutuligera Ichneumonidae Anomaloninae ?Geometridae 1 species NI Ichneumonidae Anomaloninae Geometridae Ennominae Isturgia catalaunaria Ichneumonidae Erebidae Boletobiinae Eublemma cf. reducta Ichneumonidae Campopleginae Gelechiidae 1 species NI Ichneumonidae Campopleginae Notodontidae Pygaerinae Clostera violcearia Ichneumonidae Campopleginae sp. ?Crambidae 1 species NI Ichneumonidae Campopleginae Campoplegini Charops Nymphalidae (possibly C. spinitarsis) Ichneumonidae Cremastinae ?Tineidae 1 species NI Ichneumonidae Metopiinae Erebidae Arctiinae Saenura flava Ichneumonidae Metopiinae Geometridae Sterrhinae Rhodometra sacraria Ichneumonidae Ophioninae Enicospilus sp. Lasiocampidae 1 species NI

Ichneumonidae Tryphoninae Netelia sp. Erebidae Erebinae Pandesma robusta Ichneumonidae Noctuidae Acontiinae Cardiosace perilis / Braconidae Ormyridae Ormyrus sp. Cecidosidae Scyrotis sp. * ? = Likely identification but needs further investigation; NI = Needs identification.

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Table 3 ‒ List of identified lepidopteran hosts parasitised by dipteran parasitoids.

Family Subfamily* Species* Erebidae Anobinae Plecoptera arctinotata Lymantriinae Euproctis aethiopica Lymantriinae Euproctis bicolor Lymantriinae Laelia municipalis Scoliopteryginae Anomis sp. Geometridae Ennominae Acrasia crinita Ennominae Heterostegane rectistriga Lasiocampidae Gonometinae Anadiasa jansei Lasiocampinae Eutricha obscura Noctuidae Heliothinae Helicoverpa armigera Plusiinae NI Notodontidae Thaumetopoeinae Anaphe panda Pieridae Pierinae Pontia helice helice Pyralidae NI NI Sphingidae Macroglossinae Basiothia schenki Smerinthinae Pseudoclanis postica * NI = Needs identification.

Table 4 ‒ Number of Lepidoptera-host associations recorded per region and country in the Afrotropical region (n = 11 054).

Region No. of Country No. of records records Southern Africa 10 338 South Africa 8 378 East Africa 535 Country unrecorded 1 717 Central Africa 86 Tanzania 430 Region unrecorded 67 Namibia 105 Madagascarene 25 Eswatini 103 West Africa 2 Zimbabwe 98 Arabian Peninsula 1 Kenya 96 Zambia 55 Malawi 23 Madagascar 22 Ethiopia 6 Mozambique 6 Comoro Islands 2 DRC 2 Lesotho 2 Rwanda 2 Gambia 1 Ghana 1 Mauritius 1 Oman 1 Principe 1 Sierra Leone 1 Uganda 1

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Table 5 ‒ Number of Lepidoptera-host associations (rearings) per Lepidoptera family for southern Africa (n = 11 007).

Lepidoptera family No. of rearings Lepidoptera family No. of rearings Erebidae 1 769 Somabrachyidae 20 Geometridae 1 621 Uraniidae 15 Lycaenidae 1 063 Galacticidae 13 Noctuidae 972 Bucculatricidae 12 Nymphalidae 852 Choreutidae 12 Saturniidae 520 Anomoeotidae 9 Sphingidae 511 Drepanidae 9 Lasiocampidae 486 Elachistidae 7 Hesperiidae 339 Sesiidae 7 Pieridae 286 Hepialidae 6 Crambidae 296 Hyblaeidae 6 Notodontidae 246 Lyonetiidae 5 Tortricidae 236 Brahmaeidae 4 Pyralidae 234 Cosmopterigidae 4 Gelechiidae 160 Lacturidae 4 Gracillariidae 159 Carposinidae 3 Nolidae 149 Heliozelidae 3 Papilionidae 137 Lecithoceridae 3 Limacodidae 92 Acrolepiidae 2 Nepticulidae 89 Adelidae 2 Psychidae 83 Epipyropidae 2 Eupterotidae 76 Scythrididae 2 Euteliidae 70 Bedelliidae 1 Gelechioidea 57 Blastobasidae 1 (unspecified) Zygaenidae 45 Brachodidae 1 Oecophoridae 33 Cecidosidae 1 Yponomeutidae 33 Himantopteridae 1 Tineidae 32 Incurvariidae 1 Pterophoridae 31 Micropterigidae 1 Ethmiidae 28 Prodidactidae 1 Bombycidae 24 Stathmopodidae 1 Cossidae 24 Metarbelidae 24 Plutellidae 24 Thyrididae 23 Family unrecorded 24

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Table 7 ‒ The 20 most reared Lepidoptera species (n = 930 rearings).

Number of host Number of Lepidoptera No. of Lepidoptera species species host families family rearings recorded recorded Helicoverpa armigera Noctuidae 175 136 39 Nudaurelia cytherea Saturniidae 65 60 17 Chiasmia brongusaria Geometridae 56 7 1 Achaea lienardi Erebidae 52 46 20 Eutricha capensis Lasiocampidae 44 32 13 Spodoptera littoralis Noctuidae 44 38 18 Hippotion celerio Sphingidae 42 26 7 Ascotis reciprocaria Geometridae 41 36 24 Cabera strigata Geometridae 41 2 1 Acherontia atropos Sphingidae 40 31 12 Papilio demodocus Papilionidae 37 23 5 Hypocala rostrata Erebidae 35 16 4 Saenura flava Erebidae 35 25 10 Rhodogastria amasis Erebidae 34 27 16 Vanessa cardui Nymphalidae 33 29 3 Achaea catella Erebidae 32 19 9 Hesperiidae 32 28 10 Lycaenidae 32 31 9 Isturgia deerraria Geometridae 30 15 2 Menophra jansei Geometridae 30 26 21

Table 8 ‒ Number of Lepidoptera-host associations per host taxon (mostly family) for southern Africa (n = 10 981).

No. No. No. No. Host taxon Host taxon Host taxon Host taxon reared reared reared reared Fabaceae 2 122 Stilbaceae 44 Onagraceae 6 Coriariaceae 1 Asteraceae 600 Olacaceae 43 Orchidaceae 6 Cycadaceae 1 Malvaceae 564 Bignoniaceae 42 Aquifoliaceae 5 Escalloniaceae 1 Anacardiaceae 476 Menispermaceae 35 Cactaceae 5 Gesneriaceae 1 Poaceae 351 Polygonaceae 35 Cupressaceae 5 Gisekiaceae 1 Celastraceae 344 Sapotaceae 35 Dipterocarpaceae 5 Grossulariaceae 1 Rubiaceae 310 Malpighiaceae 32 Platanaceae 5 Haloragaceae 1 Combretaceae 276 Ranunculaceae 32 Smilacaceae 5 Hypoxidaceae 1 Sapindaceae 223 Chrysobalanaceae 31 Caryophyllaceae 4 Kewaceae 1 Rhamnaceae 214 Araceae 30 Dioscoreaceae 4 Kirkiaceae 1 Euphorbiaceae 205 Icacinaceae 30 Hamamelidaceae 4 Linaceae 1 Proteaceae 198 Loganiaceae 29 Lythraceae 4 Magnoliaceae 1 Capparaceae 192 Zamiaceae 29 Strelitziaceae 4 Molluginaceae 1 Lamiaceae 186 Plumbaginaceae 28 Tropaeolaceae 4 Myoporaceae 1 Apocynaceae 179 Achariaceae 27 Violaceae 4 Nephrolepidaceae 1 Ebenaceae 177 Amaryllidaceae 27 Adiantaceae 3 Nyctaginaceae 1 Moraceae 170 Cyperaceae 24 Cunoniaceae 3 Nymphaeaceae 1 Acanthaceae 143 Nyctaginaceae 24 Dennstaedtiaceae 3 Osmundaceae 1 Loranthaceae 130 Primulaceae 24 Eriospermaceae 3 Phytolaccaceae 1 Solanaceae 127 Araliaceae 23 Erythroxylaceae 3 Putranjivaceae 1 Myrtaceae 125 Fagaceae 23 Gentianaceae 3 Resedaceae 1 Vitaceae 125 Melianthaceae 23 Musaceae 3 Saxifragaceae 1 Rutaceae 109 Thymelaeaceae 23 Oliniaceae 3 Taxodiaceae 1

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Boraginaceae 107 Commelinaceae 22 Orobanchaceae 3 Vahliaceae 1 Plant family Rosaceae 105 Restionaceae 19 Papaveraceae 3 218 unspecified Phyllanthaceae 100 Apiaceae 18 Pittosporaceae 3 Cannabaceae 95 Salvadoraceae 17 Podocarpaceae 3 Meliaceae 88 Arecaceae 16 Polygalaceae 3 Salicaceae 87 Bruniaceae 16 Ptaeroxylaceae 3 Non-plant Passifloraceae 86 Myricaceae 16 Talinaceae 3 associations: Lichens Pinaceae 85 Myrsinaceae 15 Velloziaceae 3 58 (unspecified) Asparagaceae 83 Pedaliaceae 15 Cannaceae 2 Cyanobacteria 18 Insecta Brassicaceae 82 Cucurbitaceae 12 Cistaceae 2 15 (unspecified) Homoptera Zygophyllaceae 82 Lauraceae 12 Dichapetalaceae 2 11 (unspecified) Santalaceae 77 Stangeriaceae 12 Juncaceae 2 Coccidae 8 Annonaceae 76 Cleomaceae 11 Lecythidaceae 2 Lecanoromycetes 3 Verbenaceae 74 Oxalidaceae 11 Lophiocarpaceae 2 Apidae 2 Crassulaceae 71 Portulacaceae 11 Melastomataceae 2 Bryophyta 2 Ericaceae 64 Asphodelaceae 10 Monimiaceae 2 Diaspididae 2 Urticaceae 62 Balsaminaceae 10 Pteridaceae 2 Formicidae 2 Scrophulariaceae 61 Caprifoliaceae 10 Theaceae 2 Jassidae 2 Oleaceae 60 Limeaceae 10 Valerianaceae 2 Algae 1 Aizoaceae 59 Zingiberaceae 10 Agavaceae 1 Fungi (unspecified) 1 Amaranthaceae 55 Ulmaceae 9 Begoniaceae 1 Marchantiophyta 1 Burseraceae 51 Clusiaceae 8 Bryaceae 1 Nymphalidae 1 Geraniaceae 51 Iridaceae 8 Canellaceae 1 Tettigometridae 1 Liliaceae 51 Selaginaceae 8 Caricaceae 1 Ochnaceae 48 Montiniaceae 7 Casuarinaceae 1 Convolvulaceae 45 Rhizophoraceae 7 Connaraceae 1

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Table 9 ‒ The 19 most recorded host-plant species (n = 1 478 rearings).

Host species Host family Rearings Number of Number of % Lepidoptera Lepidoptera Lepidoptera species/rearings species recorded families recorded Fabaceae 356 158 22 44 Gymnosporia buxifolia Celastraceae 144 67 10 47 Ziziphus mucronata Rhamnaceae 115 48 13 42 Acacia mearnsii Fabaceae 100 86 13 86 Fabaceae 71 31 9 44 Brachystegia spiciformis Fabaceae 59 52 10 88 Osteospermum moniliferum Asteraceae 58 39 11 67 Searsia pyroides Anacardiaceae 56 32 10 57 Julbernardia globiflora Fabaceae 55 44 10 80 Grewia occidentalis Malvaceae 54 32 13 59 Ehrharta erecta Poaceae 52 36 2 69 Searsia chirindensis Anacardiaceae 49 30 9 61 Diospyros lycioides Ebenaceae 47 33 14 70 africana Cannabaceae 46 26 15 57 Combretum apiculatum Combretaceae 46 30 14 65 Combretum molle Combretaceae 44 33 12 75 Sclerocarya birrea Anacardiaceae 43 27 9 63 Protea caffra Proteaceae 42 23 10 55 Schotia brachypetala Fabaceae 41 26 12 63

Table 10 ‒ Number of rearings of Lepidoptera-host associations per host plant family for southern Africa compared to species diversity, for the 19 most utilised plant families. Plant families with exceptionally low percentages (the proportion of no. of rearings and no. of host specie recorded per host species diversity) are indicated with an asterisk (n = 6 946 rearings).

Host family No. No. host No. No. host species No. rearings/ No. host rearings species Lepidoptera in southern host species species recorded species Africa (host diversity (%) recorded/ host recorded species diversity) species diversity (%) Fabaceae 2 201 359 747 6 374 34.5 5.6 Malvaceae 536 116 191 1 359 39.4 8.5 Asteraceae* 512 212 247 3 286 15.6 6.5 Anacardiaceae 423 67 221 453 93.4 14.8 Poaceae* 357 76 168 2 673 13.4 2.8 Celastraceae 302 39 149 336 89.9 11.6 Rubiaceae 288 92 141 866 33.3 10.6 Loranthaceae 269 32 46 234 115.0 13.7 Combretaceae 268 31 152 277 96.8 11.1 Sapindaceae 235 47 100 306 76.8 15.3 Euphorbiaceae* 227 61 118 1 482 15.3 4.1 Rhamnaceae 195 23 94 265 73.6 8.7 Capparaceae 189 32 63 273 69.2 11.7 Proteaceae 183 50 89 543 33.7 9.2 Lamiaceae 169 77 93 715 23.6 10.8 Moraceae 167 34 58 282 59.2 12 Ebenaceae 162 31 89 149 108.7 20.8 Acanthaceae 136 63 53 692 19.7 9.1 Apocynaceae* 127 63 76 1 611 7.9 3.9

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Table 13 ‒ Presenting the exponential factor (A) for the most frequently used host families for Lepidoptera families having more than 100 rearing records, in southern Africa (n = 10 478 rearings).

Lepidoptera No. of Top host No. of % of rearing No. of Exponential Lepidoptera Top host family rearing family utilised rearing records for host factor species family records records for top host families (A = D/(B-D) diversity species (B) top host family (E) x E) (southern diversity family (D) (D/B*100) Africa) (southern Africa) Erebidae 1 578 Fabaceae 376 24 104 33 1 565 6 374 Geometridae 1 340 Fabaceae 355 26 87 31 1 531 6 374 Lasiocampidae 475 Fabaceae 162 34 58 30 227 6 374 Lycaenidae 1 283 Fabaceae 316 25 66 22 509 6 374 Pieridae 305 Capparaceae 149 49 23 22 64 273 Saturniidae 514 Fabaceae 145 28 51 20 81 6 374 Noctuidae 859 Asteraceae 142 17 84 17 967 3 286 Tortricidae 207 Fabaceae 42 20 56 14 298 6 374 Papilionidae 163 Rutaceae 87 53 12 14 22 488 Nymphalidae 989 Fabaceae 183 19 56 13 324 6 374 Hesperiidae 392 Malvaceae 108 28 32 12 155 1 359 Pyralidae 215 Fabaceae 44 20 42 11 630 6 374 Sphingidae 491 Rubiaceae 80 16 46 10 111 866 Hesperiidae 392 Poaceae 93 24 32 10 155 2 637 Pyralidae 215 Celastraceae 41 19 42 10 630 336 Gelechiidae 120 Fabaceae 27 23 33 10 597 6 374 Crambidae 252 Fabaceae 35 14 49 8 526 6 374 Nolidae 149 Malvaceae 39 26 23 8 159 1 359 Nolidae 149 Combretaceae 33 22 23 7 159 277 Notodontidae 234 Fabaceae 41 18 28 6 225 6 374 Gracillariidae 156 Fabaceae 22 14 35 6 175 6 374

Table 14 ‒ Recorded Lepidoptera-host associations per subfamily and tribe in the family Fabaceae (n = 2 187).

Subfamily Tribe No, of Subfamily Tribe No. of rearings rearings 1 099 Genisteae 12 Acacieae 929 Desmodieae 7 Mimoseae 109 Robinieae 7 Ingeae 61 Undetermined to tribe 4 Faboideae 653 Swartzieae 3 Phaseoleae 147 Galegeae 2 Crotalarieae 108 Hypocalypteae 2 Millettieae 98 Psoraleeae 2 Dalbergieae 64 Caesalpinioideae 425 Indigofereae 49 Detarieae 234 Trifolieae 37 Cassieae 72 Sophoreae 33 Caesalpinieae 68 Aeschynomeneae 24 Cercideae 51 Podalyrieae 21 Undetermined to subfamily 10 Sesbanieae 20 Fabeae 13

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Table 16 ‒ Recorded Lepidoptera-host associations per species in the genus Vachellia (n = 522).

Species No. of Species No. of rearings rearings Vachellia karroo 356 Vachellia xanthophloea 3 Vachellia sieberiana 35 Vachellia borleae 2 Vachellia nilotica 27 Vachellia erioloba 2 Vachellia tortilis 27 Vachellia hebeclada 2 Vachellia robusta 16 Vachellia amythethophylla 1 Vachellia kosiensis 14 Vachellia davyi 1 Vachellia gerrardii 12 Vachellia kirkii 1 Vachellia abyssinica 8 Vachellia natalitia 1 Vachellia exuvialis 8 Vachellia seyal 1 Vachellia hockii 4 Vachellia swazica 1